As human activity along coastlines increase, degradation and destruction of coastal marine ecosystems around the globe will increase at an alarming rate. In an effort to mitigate degradation and destruction of coastal marine ecosystems, artificial reefs have been used in restoration and enhancement projects. As artificial reefs are the main method of restoring diversity to a degraded area, it is important to know the mechanisms that drive marine community assembly and diversity on those reefs. Understanding community assembly patterns of foundational species, in particular, may provide insight to community assembly patterns at higher trophic levels. Subtidal macroalgae are commonly seen as foundational species in marine environments and both deterministic and stochastic processes play a role in their assembly. Environmental gradients, which are deterministic processes, play a significant role in structuring subtidal macroalgae communities. Depth, which is negatively correlated with light, is the main driver structuring subtidal macroalgal communities, however, other gradients such as water flow, and distance to a propagule source also impact their assembly. This study sought to determine which environmental gradients play a prominent role in subtidal macroalgal community assembly. To study subtidal macroalgal community assembly, 92 artificial reef units called Reef Balls were deployed east of the Ogden Point Breakwater in Victoria, BC in June 2009. Two years passed to allow for macroalgal growth and early successional processes to occur prior to sampling the communities on thirty Reef Balls via underwater collection in July 2011. Algae were sorted by genus and dry weight was measured. To determine effects of environmental gradients on community assembly light, depth, water flow, distance to the nearest Reef Ball and distance to the breakwater were measured at each Reef Ball. A redundancy analysis revealed that depth was the most significant environmental gradient shaping algae communities and had the greatest effect on upper canopy algae. Spatial plots reveal a depth and coastline zonation of algae genera comprising the canopy. While depth was found to significantly structure algae genera found in the canopy, there was a high degree of unexplained variation in the model. This suggests that unmeasured variables such as colonization and priority effects may be driving algal community structure in the lower canopy. Differences in community structure between upper and lower canopy reveal that multiple mechanisms are responsible for shaping subtidal algal communities. Further study is required to determine the importance of stochastic colonization events and priority effects. / Graduate / 0329 / vmucciar@uvic.ca

As human activity along coastlines increase, degradation and destruction of coastal marine ecosystems around the globe will increase at an alarming rate. In an effort to mitigate degradation and destruction of coastal marine ecosystems, artificial reefs have been used in restoration and enhancement projects. As artificial reefs are the main method of restoring diversity to a degraded area, it is important to know the mechanisms that drive marine community assembly and diversity on those reefs. Understanding community assembly patterns of foundational species, in particular, may provide insight to community assembly patterns at higher trophic levels. Subtidal macroalgae are commonly seen as foundational species in marine environments and both deterministic and stochastic processes play a role in their assembly. Environmental gradients, which are deterministic processes, play a significant role in structuring subtidal macroalgae communities. Depth, which is negatively correlated with light, is the main driver structuring subtidal macroalgal communities, however, other gradients such as water flow, and distance to a propagule source also impact their assembly. This study sought to determine which environmental gradients play a prominent role in subtidal macroalgal community assembly. To study subtidal macroalgal community assembly, 92 artificial reef units called Reef Balls were deployed east of the Ogden Point Breakwater in Victoria, BC in June 2009. Two years passed to allow for macroalgal growth and early successional processes to occur prior to sampling the communities on thirty Reef Balls via underwater collection in July 2011. Algae were sorted by genus and dry weight was measured. To determine effects of environmental gradients on community assembly light, depth, water flow, distance to the nearest Reef Ball and distance to the breakwater were measured at each Reef Ball. A redundancy analysis revealed that depth was the most significant environmental gradient shaping algae communities and had the greatest effect on upper canopy algae. Spatial plots reveal a depth and coastline zonation of algae genera comprising the canopy. While depth was found to significantly structure algae genera found in the canopy, there was a high degree of unexplained variation in the model. This suggests that unmeasured variables such as colonization and priority effects may be driving algal community structure in the lower canopy. Differences in community structure between upper and lower canopy reveal that multiple mechanisms are responsible for shaping subtidal algal communities. Further study is required to determine the importance of stochastic colonization events and priority effects. / Graduate / 0329 / vmucciar@uvic.ca

This study focused on an invasive riparian reed grass, Arundo donax L., a clonal plant of the family Poaceae that is widely distributed in North America. Water availability, including water taken up from the roots locally or transported from the neighboring ramets, may affect the performance of A. donax in riparian zones. The first objective was to find out how moisture gradients affected the performance of A. donax in riparian zones. I measured leaf photosynthetic rate, leaf δ13C ratio, and plant growth-related parameters across two summer growing seasons at four transects perpendicular to the water course on the Rio Grande in South Texas. The second objective was to find out whether physiological integration existed in A. donax and how resource sharing, if any, affected plant growth. A rhizome severing experiment was conducted on five paired plots to compare growth-related parameters between plots with rhizomes severed and intact at 3, 7, and 11 weeks after treatment. Heavy water (δ 2H ~1800‰) was applied on three 1-m2 area over 3 successive days and rhizome samples were collected beyond the watering zone after 5, 24, and 48 hours of last watering. At short-term scales, A. donax performance was adversely affected by both drought and inundated conditions; over longer time scales, plant performance decreased as water availability declined in general, but biomass and stem density were similar across moisture gradients. I also found evidence of physiological integration in A. donax. Water was transported through interconnected rhizomes at least 3.5 m; transport distances averaged 1.67 m. Rhizome severing stimulated higher ramet production initially but over longer periods produced shorter thinner stems with lower flood tolerance. However, after 11 weeks of re-growth, plot-level biomass was similar between plots with severed and intact rhizomes. These results suggest that performance of A. donax is affected by water availability in riparian zones; however, clonal plant plasticity, water use efficiency, and clonal integration ameliorate impacts of water stress on the performance of A. donax. Such traits enhance its resource use, which could potentially increase competitive ability rate of establishment, and extent of this invasive species in heterogeneous riparian environments.

Phytoliths are mostly formed by groundwater silica carried upwards in a plant's vascular system and consequently accumulated in different parts of the plant. After the plant's death, the phytoliths shaped in specific morphotypes are released into the soil. Phytolith assemblages found in the soil can therefore provide information on the previous vegetation cover. The use of phytoliths as a paleoecological tool is based on their high stability in a broad spectrum of environments, even in arid conditions which are usually not suitable for preservation of pollen grains and plant macro-remains. Analysis of surface phytolith assemblages linked to the parallel study of modern vegetation is offering a good opportunity to refine our knowledge of paleoecosystems of NE Africa by evaluating the interpretation value of the fossil data. Modern phytolith assemblages are usually interpreted by using their phytolith ratio to estimate the density of tree cover, the aridity, and the proportion of C3 and C4 grasses (Barboni et al. 1999). Here we show a comparison of soil phytolith assemblages on a 400 km long climatic gradient reaching from semidesert to short grass savanna. And we also show local variability in phytolith assamblages at one of localities in comparision with grass reference collection. This master...

Canyon shaped reservoirs are characteristic by specific environmental horizontal gradients, so they enable existence of several species of the Daphnia longispina complex in one water body. Due to preference of distinct environmental conditions Daphnia species occurre in the different localities. The aim of my thesis was to analyze detail taxonomical and clonal structure of Daphnia longispina group by ten microsatellite markers on longitudinal gradient and compare it between two consecutive seasons. Simultaneously I received newly discovered divergent mitochondrial lineage from Želivka reservoir. It was confirmed, that the distribution of species and their hybrids in water reservoir was non-concidental and the taxonomic spatial distribution is in two consecutive seasons relatively constant. On the contrary the spatial and temporal distribution of clones was very heterogeneous. Clonal diversity in the interspecific hybrids was lower than in the coexisting parental species. This finding supports the hypothesis, that there exist reproductive barriers between parental genomes. Most of the clones were substantively variable, but several clones that occurred in both seasons in similar localities were found. It is possible that some clones are able of overwintering in hypolimnion and in the spring...

Submitted by Jean Medeiros (jeanletras@uepb.edu.br) on 2016-02-25T17:54:47Z No. of bitstreams: 2 PDF - Paulo Sérgio Monteiro Ferreira.pdf: 2116190 bytes, checksum: a97029f785ad5bd7d572650db6aa3e29 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) / Approved for entry into archive by Secta BC (secta.csu.bc@uepb.edu.br) on 2016-03-10T15:13:43Z (GMT) No. of bitstreams: 2 PDF - Paulo Sérgio Monteiro Ferreira.pdf: 2116190 bytes, checksum: a97029f785ad5bd7d572650db6aa3e29 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) / Made available in DSpace on 2016-03-10T15:13:43Z (GMT). No. of bitstreams: 2 PDF - Paulo Sérgio Monteiro Ferreira.pdf: 2116190 bytes, checksum: a97029f785ad5bd7d572650db6aa3e29 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) Previous issue date: 2015-02-26 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / The study of how species behave in relation to environmental gradients is an established tool in the ecology, being investigated since the pioneering studies in this field. Cactaceae is established as an important constituent family of plant communities of the Brazilian semiarid, and the change in the distribution of many genres and modes of life of this family coincides with the transition between wet and dry environments. In this context, this study aimed to test the hypothesis that the structure and diversity of cactaceas communities will be differentiated in relation to different areas of the semiarid of Paraiba, because the genera and species of this family present distinct environmental preferences. To carry out the study were selected four areas (A1, A2, A3 and A4) with pluviometric precipitations decreasing (650, 550, 450 and 350 mm respectively) in the semiarid of Paraiba. We assessed a total of 3,660 individuals of Cactaceae, distributed in six species: Cereus jamacaru DC., Melocactus zehntneri (Britton & Rose) Luetzelb, Opuntia inamoena K. Schum, Pilosocereus gounellei (F. A. C. Weber) Byles & G. D. Rowley, Pilosocereus pachycladus F. Ritter and Tacinga palmadora (Britton & Rose) N.P. Taylor & Stuppy. A1 presented 196 individuals divided into three species (C. jamacaru, O. inamoena and P. pachycladus); A2 was represented by 642 individuals, A3 presented 1,447 individuals and A4 1,375 individuals divided among the six species mentioned. The cactaceas species demonstrated patterns of Importance Value Index (IVI) and Relative Volume (VR) between the different areas. A1 presented the highest individuals (mean height = 4.96 m) and with bigger DAS (mean DAS = 13,34 cm), and having the largest volume of cactaceas (21.42 m 3 ); A2 showed more diversity (H '= 1.39) and equability (J' = 0.83); A3 had the lowest values of DAS (mean DAS = 4.33 cm), height (mean height= 0.85 m), diversity (H'= 0.58), equability (J'= 0.39) and volume of cactaceas (4.92 m ); and A4 presented, in most cases, intermediate values between A2 and A3. The greatest richness and abundance of species were related to the locations with low rainfall and more clayey soils had greater diversity of cactaceas. The nMDS (non-Metric Multidimensional Scaling) produced by the environmental variables separated the samples into four groups and the cluster analysis showed differentiated configurations of these attributes within their own areas. The groups differed significantly according to the permutation of Monte Carlo test that were between the Areas 1 and 2; 1 and 4; 2 and 4; and 3 and 4. Of these environmental variables, the predictive model given by BIO-ENV (Biotic-Environmental) suggested that the percentage of clay in the soil and EVI (Enhanced Vegetation Index) are those that best explain the variations of cactaceas communities. The CCA diagram (Canonical Correspondence Analysis), produced 3 with the predicted variables by BIO-ENV showed that the cactacea species that occurring in of Paraiba semiarid have different environmental preferences, revealing that the species P. pachycladus and C. jamacaru were positively related to areas with higher indices of EVI and rainfall; M. zehntneri and P. gounellei adapted better in areas with soils with higher proportion of clay and silt; O. inamoena, was negatively related with higher proportions of medium sand; and T. palmadora, showed no patterns related to variables given by the BIOENV. The significant correlations between the axes of the CCA and the attributes of the structure of the species (abundance, height and DAS), were some times moderate (r 0.40 to 0.69) and most of the times stronger (r ≥ 0.70) and the species that presented best linear associations with the variables tested were M. zehntneri, P. gounellei and P. pachycladus. / O estudo de como as espécies se comportam em relação aos gradientes ambientais é uma ferramenta estabelecida na ecologia, sendo investigado desde os estudos pioneiros nesse campo. Cactaceae, estabelece-se como uma importante família constituinte das comunidades vegetais do semiárido brasileiro, e a mudança na distribuição de muitos gêneros e formas de vida dessa família coincide com a transição entre ambientes secos e úmidos. Nesse contexto, o presente estudo buscou testar a hipótese de que a estrutura e diversidade das comunidades de cactáceas serão diferenciadas em relação às diferentes áreas do semiárido paraibano, pelo fato dos gêneros e espécies dessa família apresentarem preferências ambientais distintas. Para a realização do estudo foram selecionadas quatro áreas (A1, A2, A3 e A4) com precipitações pluviométricas decrescentes (650, 550, 450 e 350 mm respectivamente) no semiárido paraibano. Foram aferidos ao todo 3.660 indivíduos de Cactaceae, distribuídos em seis espécies: Cereus jamacaru DC., Melocactus zehntneri (Britton & Rose) Luetzelb, Opuntia inamoena K. Schum, Pilosocereus gounellei (F. A. C. Weber) Byles & G. D. Rowley, Pilosocereus pachycladus F. Ritter e Tacinga palmadora (Britton & Rose) N.P. Taylor & Stuppy. A1 apresentou 196 indivíduos divididos em três espécies (C. jamacaru, O. inamoena e P. pachycladus); A2 foi representada por 642 indivíduos, A3 apresentou 1.447 indivíduos e A4 1.375 indivíduos divididos entre as seis espécies citadas. As espécies de cactáceas demonstraram padrões de Índice de Valor de Importância (IVI) e Volume Relativo (VR) distintos entre as Áreas. A1 apresentou os indivíduos mais altos (Alt. med. = 4,96 m) e de maior DAS (DAS med. = 13,34 cm), possuindo também o maior volume de cactáceas (21,42 m 3 ); a A2 demonstrou maior diversidade (H’ = 1,39) e equabilidade (J’ = 0,83); A3 apresentou os menores valores de DAS (DAS med. = 4,33 cm), altura (Alt. med.= 0,85 m), diversidade (H’ = 0,58), equabilidade (J’ = 0,39) e volume de cactáceas (4,92 m ); já A4 apresentou, na maioria das vezes, valores intermediários entre A2 e A3. As maiores riquezas e abundâncias estiveram relacionadas aos locais de menores precipitações e os solos mais argilosos foram os que comportaram maior diversidade de cactáceas. O nMDS (Escalonamento Multidimensional não-Métrico) produzido através das variáveis ambientais separou as amostras em quatro grupos e a análise de agrupamento demonstrou configurações diferenciadas desses atributos dentro das próprias Áreas. Os grupos que diferiram estatisticamente segundo a permutação do teste de Monte Carlo foram entre as Áreas 1 e 2; 1 e 4; 2 e 4; e 3 e 4. Destas variáveis ambientais, o modelo preditivo dado pelo BIO-ENV (Biotic-Environmental) sugeriu que a % de Argila no solo e o EVI (Índice de Vegetação 3 Aprimorada) são as que melhor explicam a variação das comunidades de cactáceas. O diagrama de CCA (Análise de Correspondência Canônica), produzido com as variáveis preditas pelo BIO-ENV, demonstrou que as espécies de cactáceas ocorrentes no semiárido paraibano apresentam preferências ambientais distintas, revelando que as espécies P. pachycladus e C. jamacaru foram positivamente relacionadas às áreas com maiores índices de EVI e Precipitação; M. zehntneri e P. gounellei, foram melhor adaptadas a habitarem em ambientes com solos com maior proporção de Argila e Silte; O. inamoena, esteve negativamente relacionada às maiores proporções de Areia média; e T. palmadora, não demonstrou padrões relacionados as variáveis dadas pelo BIO-ENV. As correlações significativas entre os eixos da CCA e os atributos da estrutura das espécies (abundância, altura e DAS), foram algumas vezes moderadas (r 0,40-0,69) e na maioria das vezes fortes (r ≥ 0,70) e as espécies que apresentaram melhores associações lineares com as variáveis testadas foram M. zehntneri, P. gounellei e P. pachycladus.

Tropical forests of the world are fast disappearing and there is a race to understand patterns of species distribution in space and time. Studying species distributions can provide better frameworks for conservation of these ecologically important patches of floral and faunal diversity. The island of Sri Lanka is a well known harbour of unique and highly threatened biodiversity. Tropical lowland forest is remnant in the south-west of the island now mainly existing in small patches. While most are small disturbed fragments, Sinharaja Forest Reserve represents one of the largest remaining patches of this important ecosystem. As a UNESCO World Heritage Site and a Man and Biosphere Reserve, it has a dual role as a conservation area and a historically important resource forest. While the distribution of vegetation diversity has been well documented, analyses of invertebrate species distributions are lacking. This thesis investigated a key arthropod group, ground dwelling ants, in relation to environmental gradients within the forest. Cumulative results demonstrate the high diversity of the forest patch. In an area representing less than half the reserve, over 173 ground dwelling ant species were found in distinct assemblages throughout the forest. Since the forest is located upon a series of parallel ridges, ant species distribution was first analysed in terms of this small elevation change. Species richness declined over a vertical incline from 430 m to 660 m, highlighting a possible small-scale, mountain mass effect. This section of the reserve is also characterised by a patch of once-logged forest (30 years previously). A study was undertaken to investigate whether there were residual effects of selective logging on the reserve. / Significant differences between species assemblages in once-logged forest and unlogged forest add to growing evidence that selectively logged forests continue to remain distinct from unlogged forest even after decades of regeneration. Ant distribution was then analysed for their relationship with habitat heterogeneity and tree species distribution. Long-term research on tree species in the SFR has demonstrated a close relationship to habitat complexity. Ant species appear to respond more to the structural heterogeneity of the vegetation than to actual topographic variation within the forest. From a conservation perspective, maintaining the integrity of this highly diverse forest is imperative. The impact of anthropogenic land uses surrounding the forest was investigated in terms of ant assemblages along the forest edges. Significant differences were found between assemblages within the edges bordered by different matrix types. Even relatively large forest remnants can be affected by the surrounding matrix land uses and encouraging the growth of structurally similar vegetation and maintaining low disturbance along the borders should attenuate the effect of the edge. Overall, the highly heterogeneous distribution of ant assemblages within the SFR demonstrates the potential for other small patches to be harbours of further species diversity. Future research should be undertaken to assess the diversity and distribution of ant species within this region and encourage the protection of this remnant diversity.

Understanding what drives variation in species diversity in space and time and limits coexistence in local communities is a main focus of community ecology and biogeography. My doctoral work aims to document patterns of ant diversity and explore the possible ecological mechanisms leading to these patterns. Elucidating the processes by which communities assemble and species coexist might help explain spatial variation in species diversity. Using a combination of manipulative experiments, broad-scale surveys, behavioral assays and phylogenetic analyses, I examine which ecological processes account for the number of species coexisting in ant communities. Ants are found in most terrestrial habitats, where they are abundant, diverse and easy to sample (Agosti et al. 2000). Hölldobler and Wilson (1990) noted that competition was the hallmark of ant ecology, and we know that ant diversity varies along environmental gradients (Kusnezov 1957). Thus ants are an ideal taxon to examine the factors shaping the structure of ecological communities and how the determinants of community structure vary in space.

Shallow-water hydrothermal vents occur world-wide in regions of volcanic activity. The vents located at Tutum Bay, Ambitle Island, Papua New Guinea are unique in that the vent fluids and surrounding sediments contain some of the highest concentrations of arsenic in a natural system. This study addresses the effects of the vent system on the benthic communities, focusing on the eukaryotes, macrofauna, meiofauna and bacteria. Samples were collected in November 2003 and May/June 2005. Analysis of the 2003 macrofaunal samples indicated that pH, rather than arsenic was influencing the benthic community, and that the hydrothermal influence occurred at a greater distance than expected. Results of more intensive sampling carried out in 2005 are the primary focus of this dissertation. The pore water and sediment characteristics revealed distinct physical habitats corresponding with distance from the vent. There was a trend of decreasing temperature and arsenic concentration and increasing salinity and pH with distance from the vent. The vent sediment was poorly sorted volcanic gravel, while sediments along the transect showed a gradient from fine, well sorted volcanic sands to coarser carbonate sands farther away. The macrofauna showed a trend of increasing diversity with distance from the vent and similar taxa were present in both the 2003 and 2005 samples. The vent community was dominated by the polychaete Capitella cf. capitata. The inner transect from 30 m to 140 m had low diversity. Dominant taxa included thalassinid shrimp and the amphipod Platyischnopus sp.A. The 180 m to 300 m sites had significantly higher diversity. The Danlum Bay reference site had relatively higher diversity than the nearshore transect sites and was dominated by deposit feeding polychaetes. Macrofaunal community structure was influenced by the sediment characteristics, notably by CaCO3 content, sorting and median grain size. The meiofaunal community also showed changes with distance from the vent. Chromadorid nematodes were dominant at the vent site and were a major component of the meiofauna at most sites, along with copepods. The meiofaunal community at the reference site showed greater similarity to the vent community and both sites had low abundances. Nematodes were more abundant than copepods near the vent, but copepods were more abundant farther offshore and at the reference site. Meiofaunal community structure was influenced primarily by the pore water temperature and salinity. Biological interactions with the macrofaunal community through physical disturbance and predation may also influence the meiofaunal community.  The molecular analysis of eukaryotic and bacterial diversity also revealed changes with distance from the vent. The 0 m and reference sites grouped together due to the presence of fungal sequences and the 140 m and 300 m sites grouped together due to a common molluscan sequence. Metazoans and fungi dominated the eukaryote sequences. The most abundant eukaryotic OTUs included fungi matching Paecilomyces sp. and Cladosporium cladosporioides and metazoans matching Viscosia viscosa (Nematoda) and Astarte castanea represented by 24 phyla and was dominated by Actinobacteria and γ-Proteobacteria. More bacterial phyla were present near the vent, while more overall OTUs were found at the intermediate sites along the transect. The most distant site had much lower diversity dominated by Firmicutes. The macrofaunal community had the strongest correlation with environmental variables. Comparison between the meiofauna and the metazoan sequences showed the proportion of nematodes found in both datasets were comparable, but the meiofauna analysis found a higher proportion of arthropods, while the molecular results were disproportionally high for platyhelminthes. Overall, the vents increased the complexity of the system by creating unique habitats. The extreme environment created by the hydrothermal activity maintained the surrounding habitat at an early successional stage colonized by a few opportunistic species. There was a gradation in the benthic communities away from the vent towards a more carbonate based climax community. The low pH environment had an effect on the sediment composition, which in turn influenced the benthic community. These findings can serve as a model for studying the potential effects of ocean acidification and climate change on benthic communities and marine biocomplexity.

The goal of this dissertation is to further our understanding of how spatially heterogeneous landscapes may impact the formation of range boundaries that then aggregate to form large-scale biodiversity patterns. These patterns have been analyzed from many different perspectives by ecologists, evolutionary biologist, and physiologists using a variety of different theoretical, statistical, and mechanistic models. For some species, there is an obvious abrupt change in the environment causing a range boundary. Other environments change gradually, and it is unclear why species fail to adapt and expand their range. The first chapter develops a novel theoretical model of how the establishment of new mutations allows for adaptation to an environmental gradient, when there is no genetic variation for the trait that limits the range. Shallow environmental gradients favor mutations that arise nearer to the range margin, have smaller phenotypic effects, and allow for proportionately larger expansions than steep gradients. Mutations that allow for range expansion tend to have large phenotypic effects causing substantial range expansions. Spatial and temporal variation in climatic and environmental variables is important for understanding species response to climate change. The second chapter uses a mechanistic model to simulate switchgrass (Panicum virgatum L.) productivity across the central and eastern U.S. for current and future climate conditions. Florida and the Gulf Coast of Texas and Louisiana have the highest predicted current and future yields. Regions where future temperature and precipitation are anticipated to increase, larger future yields are expected. Large-scale geographic patterns of biodiversity are documented for many taxa. The mechanisms allowing for the coexistence of more of species in certain regions are poorly understood. The third chapter employs a newly developed wavelet lifting technique to extract scale-dependent patterns from irregularly spaced two-dimensional ecological data and analyzes the relationship between breeding avian richness and four energy variables. Evapotranspiration, temperature, and precipitation are significant predictors of richness at intermediate-to-large scales. Net primary production is the only significant predictor across small-to-large scales, and explains the most variation in richness (~40%) at an intermediate scale. Changes in the species-energy relationship with scale, may indicate a shift in the mechanism governing species richness. / text